Alison F. Richard

Evolution in the hypervariable environment of Madagascar

We show that the diverse ecoregions of Madagascar share one distinctive climatic feature: unpredictable intra- or interannual precipitation compared with other regions with comparable rainfall. Climatic unpredictability is associated with unpredictable patterns of fruiting and flowering. It is argued that these features have shaped the evolution of distinctive characteristics in the mammalian fauna of the island. Endemic Herpestidae and Tenrecidae and members of five endemic primate families differ from closely related species elsewhere, exhibiting extremes of “fastness” and “slowness” in their life histories. Climatic features may also account for the dearth of frugivorous birds and mammals in Madagascar, and for the evolutionary prevalence of species with large body mass.

Weed macaques: The evolutionary implications of macaque feeding ecology

Patterns of feeding ecology among the living macaques conform poorly with recognized phyletic distinctions within the genus because there is an important ecological division which cross-cuts phyletic groupings. This division, between weed species and non-weed species, is based on the differing abilities of macaques to tolerate and even prosper in close association with human settlements. Based on available information about their ecology in the wild, we tentatively assign macaque species to these two categories. Finally, we consider the implications of our argument for scenarios of the initial spread of the macaques.

Life in the slow lane? Demography and life histories of male and female sifaka ( Propithecus verreauxi verreauxi )

A 16-year study of wild, unprovisioned sifaka Propithecus verreauxi verreauxi at Beza Mahafaly in south-west Madagascar provides estimates of age-specific fertility, mortality, and dispersal in a population of 426 marked animals, and longitudinal records of individual life histories. Sifaka females give birth for the first time later and live longer, for their size, than mammals in other orders; they also give birth later and continue reproducing longer, for their size, than other primates. Theory postulates that these features, commonly referred to as bet-hedging, evolve in unpredictable environments in association with widely varying infant survival and a trade-off between reproductive effort and adult survival. The climate of south-west Madagascar is highly unpredictable compared to almost all other regions in the tropics with similar average rainfall, and we argue that sifaka females are bet-hedgers par excellence. Male sifaka, in contrast, become reproductively active at an earlier age than females, and are less likely to have long lives than females. The atypical direction of this asymmetry between males and females reflects a ‘slowing down’ of female life histories rather than a ‘speeding up’ of male life histories. Two other unusual features of sifaka biology and behaviour may be linked to the unpredictability of Madagascars climate: intense local competition between females, and a sex ratio at birth strongly biased in favour of males in most years. In drought years, reproductive females must cope with suddenly intensified resource constraints. This, in turn, may strongly limit the number of ‘breeding slots’ available over the long-term for females.

Intra-specific variation in the social organization and ecology of Propithecus verreauxi.

This paper presents information collected during an 18-month study of four groups of Propithecus verreauxi living in the north-west and south of Madagascar. Various aspects of the s

Reproduction in Free-Ranging Male Propithecus verreauxi: The Hormonal Correlates of Mating and Aggression

Endocrine studies of captive strepsirrhine primates suggest that physical environment and social factors mediate inter-individual variations in testicular function and serum testosterone (sT) in males. While these studies have made major contributions to our understanding of the individual proximate mechanisms influencing androgen activity in male strepsirrhines, none have investigated how these mechanisms work coincidentally in free-ranging populations. In this study we used fecal steroid analysis to examine androgen-behavior interactions associated with reproduction in free-ranging male Propithecus verreauxi. Behavioral and hormone data were collected from two social groups during the 1990-91 and 1991-92 breeding seasons at Beza Mahafaly, Madagascar. Solid phase and radioimmunoassay techniques were used to quantify testosterone (T) in 105 desiccated fecal samples collected weekly from seven males. Results suggest that 1) solid phase extraction and radioimmunoassay techniques were reliable and accurate methods for quantifying T in sifaka feces; 2) fecal T (fT) elevations spanned a minimum of 4 months, peak levels occurring 1 month prior to the January onset of the breeding season; 3) fecal T concentrations were influenced by developmental factors and, among mature males, social factors associated with rank, intergroup aggression, and group instability.

Aggressive competition between males, female-controlled polygyny and sexual monomorphism in a Malagasy primate, Propithecus verreauxi

Abstract The evolutionary determinants of sexual dimorphism among primates continue to be debated. This paper uses field observations of a Malagasy primate, Propithecus verreauxi, to argue that one problem underlying this debates is the unrecognized heterogeneity of mating system categories used in comparative analyses investigating the importance of sexual selection in determining sexual dimorphism. It is suggested that at least one new mating system category warrants recognition, that of female-controlled polygyny, with predictions for sexual dimorphism that are in contrast to those of male-controlled polygyny.

Ecology of rhesus macaques (Macaca mulatta) in northwest Pakistan

The feeding ecology of rhesus monkeys,Macaca mulatta, was studied between 1978 and 1981. The study site, located in the Murree Hills of northwestern Pakistan, supported a mixed coniferous-deciduous forest community and was characterized by a high degree of human disturbance. We used a linear transect method to sample the species composition and structure of the vegetation. Comparison of these data with historical records showed that the forest has undergone major changes in the last hundred years. Data on feeding behavior were collected through on-the-minute focal animal sampling. The monkeys spent about 45% of the day feeding. Their preferred foods were grass, clover, and other ground herbs that occur in the disturbed sites. Fruit accounted for less than 9% of feeding records. The rhesus monkey may be pre-adapted to living in disturbed-site, forest-edge communities. The evolutionary history of Macaca mulatta may be tied closely to the disappearance of forest and the spread of meadows and savannahs over the last million years.

Birth season testosterone levels in male Verreaux’s sifaka, Propithecus verreauxi: insights into socio-demographic factors mediating seasonal testicular function

Abstract Previous research suggests that although reproduction and testicular function in wild sifaka (Propithecus verreauxi) are highly seasonal, birth season elevations in fecal testosterone (T) in transferring males indicate that social factors may be as important as climatic factors in regulating reproductive function in sifaka. This paper examines the relationship of male dispersal and social status to the patterning of birth season aggression and testicular function in P. verreauxi at Beza Mahafaly Special Reserve, Madagascar. Behavioral and hormonal data were collected from a total of 38 adult males, 15 residing in seven stable groups and 23 living in eight unstable groups, yielding 186 fecal samples and 493 focal animal hours of observation. The results suggest that birth season elevations in fecal testosterone are the consequence of social disruption resulting from male movements between groups and the particular responses of individual males to dispersal events. Hormonal responses to dispersal events appear to precede behavioral responses and occasionally register reactions not concomitantly evident in the behavioral response, and may be predictive of future events. Hormonal reactions occurred primarily in resident alpha males, suggesting that they differ in their assessment of destabilizing influences, perhaps due to different reproductive opportunities and/or investment. These findings are important for the new insights they provide into the role of androgens in mediating male dispersal, life history, and reproductive strategies, and suggest that investigations of androgen-behavior interactions in free-ranging populations can be a powerful new tool for assessing the contextual and motivational basis of social behavior.

Social boundaries in a Malagasy Prosimian, the Sifaka (Propithecus verreauxi)

Using focal animal samples, the social organization of sifakas was studied in two forests for 2500 hr spread over 18 months. Data were also obtained on the size and composition of groups at two other sites. The size and adult sex ratio of groups varied widely within populations, although population-wide sex ratios approached unity. During the brief annual mating season, some males mated with females belonging to other groups. The response of both male and female group members to the approach of a nongroup male appeared to depend on his identity. It is argued that while group size may be constrained by the availability of resources, group composition reflects the effects of small-sample demographic variations rather than the outcome of reproductive competition among or between males and females. Spatial boundaries of groups do not coincide with social or reproductive boundaries even within a single breeding season; further study is needed of the processes whereby relationships are established and maintained between individuals from different groups.

Genetic population structure of the white sifaka ( Propithecus verreauxi verreauxi ) at Beza Mahafaly Special Reserve, southwest Madagascar (1992-2001)

Gene flow within and between social groups is contingent on behaviourally mediated patterns of mating and dispersal. To understand how these patterns affect the genetic structure of primate populations, long‐term data are required. In this study, we analyse 10 years of demographic and genetic data from a wild lemur population (Propithecus verreauxi verreauxi) at Beza Mahafaly Special Reserve, southwest Madagascar. Our goal is to specify how patterns of mating and dispersal determine kinship and genetic diversity among animals in the population. Specifically, we use microsatellite, parentage, and census data to obtain estimates of genetic subdivision (FST), within group homozygosity (FIS), and relatedness (r) within and among social groups in the population. We analyse different classes of individuals (i.e. adults, offspring, males, females) separately in order to discern which classes most strongly influence aspects of population structure. Microsatellite data reveal that, across years, offspring are consistently more heterozygous than expected within social groups (FIS mean = −0.068) while adults show both positive and negative deviations from expected genotypic frequencies within groups (FIS mean = 0.003). Offspring cohorts are more genetically subdivided than adults (FST mean = 0.108 vs. 0.052) and adult females are more genetically subdivided than adult males (FST mean = 0.098 vs. 0.046). As the proportion of females in social groups increases, the proportion of offspring sired by resident males decreases. Offspring are characterized by a heterozygote excess as resident males (vs. nonresident males) sire the majority of offspring within groups. We link these genetic data to patterns of female philopatry, male dispersal, exogamy, and offspring sex‐ratio. Overall, these data reveal how mating and dispersal tactics influence the genetic population structure in this species.